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Selection and propagation of superior teak for quality improvement in plantations : case study of the ICSB/Cirad-Forêt joint project

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Quality Timber Products of

Teak

from Sustainable Forest Management

ProœedmssoftheInremationalConkrcrne:-tQualityT~ProdIlct5ofTrUfromSustaINbk-Fon>stManagement Peechl.lndia. 2-5Deœmber2D.D

Edilors

K.M. Bhat, K. K. N. Nair, K. V. Bhat

E.M. Muralidharan and

J.

K. Sharma

Kerala Forest Research Institute, Peechi 680 653, lndia

AnInstitution01KeraaStateCoociIforScienœ.T«hrdogyandEnvironrnmt(KSCSTE)

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02005 Keub Forr'lt Rr'lurch IMljtute

The Kerala Forest Research Instltute (KFRI)isone of theSIXinstitutions under

theKenlOiStateCounol forSoenœ, Technology iU1d Ennronmenl (KSCSTE)of the Go.'emment of Kerala, estOlbhshed in 1975. Byronducting time-bound mulbdlSCIpllnary applled research ln lhrustareasof tropIcalforestry,KFRlhas crt'itecllimchr .. mongthele..dIng fores!research orgamutions in the IropICl. Th", Institut'" unaertakes mulbd1sclplmaryresearch on ..Il asprctsoftroptcal

fo~trymduding ...ood samœ and technotogy, wtIdlile biologyand

socio-fCOnOIJUC5undrrtheProgra1ll/Tll!'DiVlSlOOS.KFRIhas a Sub-Centn ..tNi1.l.mbur andliFieldSUbon atVrl~forcalT}'mg out nursery andpbntabonbuis, grrmplasm roli«bon,~c.KFRItwthe1argestroUl!'ctJonofbamboo and ratbl sprrirsinInch.a fOI" rrsearm andconsernbon~AlsoilNi1ambur,lhere isilTnk Museum. theonIyoneclltskJnd.~'Otedtoa singletnorsprciI!'Sinthe wOI"Id,Itisopmlopubbc,researchers.fores!offioalsandethersmlen'stedin

""

InitmitiouiTropinl1im~rOrpniution

TheIntermtK'NI Tl'Of'ICI'l TunbeOrpniuIion.(ITIO)isaninll!'lgO\wnmmtal orgmiubonwttid\bnngslogdhrf oountnesiNtprodl.lCl!'andconsumr tropical biTIbft- todl5CUS$andexdungeinformationanddeveloppoIiorsonail~

cl the woOdtroptaIbmbrr~y.TherrroishudquarteredmYokohama.

Jilpanandlnluly2001hadSlmrmbrrs.induding theEuropean Commuruty,

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logether rtpresell95f'l"I"œntof ...oridtrMte intropial.~and 75 perC'l!'ntoflhewotld'slropeal forests.Therrronorognizeslhat01 conllnuing supPIyoflroplCillllmber on the world markddependson quahtyinformlllion.

about thelTadeand marketplaœ,on ef6orn1 bmbrrproduclion.andproœs5U'lg methods and onsusbmllbieforest Iniln.J.grmrnl practices

Publishtd by:

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Tri: 91-487-2699037, 2699061-64;Fax:91-487-26992..19 Email:UnOkfnC'rS Website: hllp://www.kfri.org

...

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ISBN81 -85().l1-63-6

KFRI LibraryCataloguing-in-Publication Daia

QUAlITY TIMBER l'RODUCTS OF TEAK FROM SUSTINABLE FOREST MANAGEMNET

I,Teakproducts 2.Sustainableforestmanagement-teak 3. TImber œrbfinlion 4 Tt'("IoruigrandIS

1.Bhal.K.M.IIKeralaFûffstResearch in.<titulr 1Il.lnlemab(Xl&1 Tropical TImberOrganiulion

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Typeset: KFRl

Printt'd&tLUll\l('ff,lhnssur6lltJU20,IndU.

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Selection and Propagation of Superior Teak for

Quality Improvement in Plantations: case study of

the ICSB/Cirad-forêt joint project

D. K. S. Cohl, D. AlJoysiusl,}. Gidiman ', H. H. Chanl, B. MalleFand O. Monteuuis2

llMoprise'Corporoltion Sdn Bhd., P. O.Box60793, 91017 Tawau, Sabah, Malaysia

E·mail: [email protected];dil\·K1@[email protected]; jgidiman@holmaiLrom; chanhh@pl",ringmr 'CIRAD-Foret, TA IOIC, 8.JilIarguet. J.l398 Montpellier. Cedex 5. France

E·mail: mallet@cirad-fr; [email protected]

ABSTRACf

Innoprise Corporation Sdn Bhd (JCSS). an in\'estment subsidiary of the Sabah Foundation, ha!> embartœdon an e:densh-{' Research and lÀ'velopmenl program for leak(TtdotUJSfI",dlS),ahighly prized limbe!species. Through ICSB's joinl collaboriltion with CIRAD-Foriot. a French R and D ocganization., a plant improvemenl program wasiniti.lledinthe early 1990'$. Specialeffortsha...e been devoted 10 vegetative propagation strategiesb.lsedonthede'l'elopment al efficleJlt lechniqlll.'S.

bothal the laboralory and al the nwseryIeo.'els.~Ialerialsfrommature selected plus ltees (ortels)from

ilbroad genelic base,and seeds of preo>uD"lt"d rugh genelic value bul in reslricled number, were multiplied byusingilweU-deveJoped tissue culture techruque al thePIaniBtotechnology uboratory. Additionally, seed lois from natural foresl stands, pl.mtationsilnda mulli-provenanceclonaiseed orchard(CSO)wereobtained.TheseWeil!germinatedandused 10 establish two main prO'.enancel

progenytrialsin two dlfft'reflt locations within ICSBconœssion in easl Sabah.Malaysia,in addition to olhernuml'rou.~conservation and demonstralion pIOIS. Data of six-year old trees from these

pron'nance/pl'Ob~ny triab ilIustrated the great potenlial of theCS(}materials esperi.1Uy in terms of

J;l1Jwlh rate rompared 10 the other origins under the pre\'aiting conditions. Advanœd selection. of J;('fM)lypes rombining v.lrious traits of major ('((Inomical valueie.wood charaderistics,isunderway. These genolypes andtheirprogenies willhefurther testedindifferent sitesintheimmed~lte future. The ownership of such arichgermplasm coupled 10 the effJCiency of the develol"-'d techniques offers

i\ tremendous potential for genetic improvement, selection and mass production for large-scale plant.ltions of higher yield and superior quality. COll1JTl('m.ll production of sclcct('(i nmleriills from this rich genetic base will !hm ensue as tissue culture-issued materials for overseas markets. Ktywords: clonaI deployment, field trials, genetic resources. germplilsm, provenilnce/progeny

trial, superior planling stock production, vegetative propagillion, tissue culture.

INTROOUcnON

Teak was firsl introduœd in Kota Marudu in the state of Sabah, Malaysia, by the Dulch TobaccoCompany in the19205',but did not bccome popular until the early lC)lX)s(Lapongan, 1998). Since then, private planlation owners and oil palm companies have grown teak mainly by intercropping with oil palm orothercashcrops, with parlicular menlion forthe mosaic IOxIOm inlerplanted with IOxIOm oil palm desIgnexperimented olt Balung Plantation in Sabah

(Salleh, 1995; White, 2002). Howevcr, the teolk planting material uscdWolSnot spccificallysclc<:ted for such plantation systems.

o.·er time, lackof informaIionon lheoptimal density and spacinginan inlercropping system and thcreby, fcarof potentiallœses 10 theyieldof associatedcrops, resultedinthe planling of tcak mainly along borders of many oil palm plantations and to sorne extent, roadsides. Moroo\'cr, the reputation of long rotation

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furthcr in Sabah and Malaysia asa whole. Asa result, investors nowadaysare morecagerto invcst in theoil palm industry due to the more lucrative retums thal

tanbeobtained in a timeperiod asshort as threeyears.

Planl~wereno!.evCll encouraged by thefinding from

a cost analysis of teak planling in Malaysia that

estimaled Icakrould be viable as a plantationaopC\'en

wilh an increase of20%in the rost of planting and a drop in prieeona 15-year rotation (Krishnapillayd

al., 1997). Nevertheless, while the interest in teak planting within Malaysia declines in the face of competition

nom

othercashcropssuch as cocoa,riœ,

rubber and especially oil palm for whkh the country hasfo,y...gamoo theworldproductionleadefship, the scenarioisjustthe opposite inother partsofthe

world.

Countriesinthe tropical and sub-Iropical zones such as Central and South America, Arrica, India, Australia, and other

parts

of South-eastAsiabegin embarking on large-scaled planting of teak, recognizing it as a prime candidate for forest plantationestablishmenl in termsof valueand world market demand. Accordingly, partiaJiarattentionis

being devoled to means for improving quantitatively and qualitalively the yields in shorter time frames than tradilionally expeded. Since the early 1990'5, Innoprise Corporation. the investment subsidiary of the Sabah Foundation, a stale-run organization. and CIRAD-Forêt, a French Researçh and J:)evelopment Organization havebeenworking together, aiming at this ob;ective.

CONSTITITrING GENFrlCAllY-RICH BASE POPULATIONS

Securing Ihe access to a genetically broad-based population is an assel of magnitude importance for developing simple or more sophisticated Iree improvement strategies. This is particularly true for leak considering Ihe noticeable differenœs among provenances, progenies and even half-sib (Bath, 2000). Relevantly, one of our priorilies hasbeento galher as many leak origins as possible and 10 establish properly designed base population plots within ICSB'sconœssions.

The first batchof seedsoriginating from the Solomon island seed-sourœ was introduced in 1989.Asfor

manyseed sources, information on the accu rate natural origin ofthis plant malerial isIacking,although Tenasserim (Myanmar, ex Bunna) seems 10 be the mosl Iikely provenance (Kevin White, personal communication).Theresultingseedlingswcre planled bythe Planllmprovem<nland SœdProdUctiOll(l'lSP forshort) UJÙt asademonstrational plot in theLuasong Foreslry Center, located aboul two hour drive from Tawau,oo theeaslcoastof Sabah. This planl material

hasbeen thriving sÎnœ lhen, showing remarkable perfonnanœs under local conditions with average diameter and height ol 2·3cmand4 m inthe firsl2-3 yeats resp«tively,followedby •gradu" evening 001 inaverageheightof2to3minsubsequenl years.The trees give rise10 long c1ear boles with delayed llowering, bearing inmind that in teak.the tater the altainment of the flowering stage, the longerthe boIe and therefore, the higherthe value.

Thesecondintroduction ofseeds was made in 1995, thislimeroII<ctedfrornthe I..ksœd sbndbeIooging

tathe ForestResearchlnstitute of Malaysia in Perlîs, Iocated in the north of Peninsular Malaysia close 10 theborderwithThailand.Thereisstill abasicneed 10 docummtlheaccurateprovenanœsoftheseseeds.The

smalJ amount ofseedsfromthisbalch wasgerminated under invitroconditions at the Plant Biotechnology Laboratory (PBL) following the proœduredescribed by Monteuuisd al. (1998), and then planted as yet anotherdemonstrational plot in Luasong.

ln 1996, in order 10 wÎden the existing genetic baseof teak in our project, ICSB jointly with Cirad-Foret procuredseedsfrom two extensive sources - from natural foreststandsor plantationsand fromprogenies produced from a multi-provenance clonaI seed orçhard in Ivory Coast. Altogelher, there were 77 seedlots comprised of India, Thailand, Papua New Guinea, Tanzania, Ivory Coast, Salomon Island, Indonesia, Segama (Sabah) and Perlis (Peninsular Mli<lysœ)(f.bles hnd2).TheseedsweregenniNled either at the PISP nursery in luasong or at the PBL, particularly those with presumably low gennination capacity. Adetailed report on theseedling procedure ., the PBl hasbeen published (Monleuu;set<li., 1998). However, in the same way as for many species introduœd as exotics, tracing the accurate natural origins of every seed-sourœ as weil as the number

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Table 1. List orthe \'<lriolls ...--edlotsobtained,germinated and pJantcd within our proJcct

Pro\'t'n·lOCt...; India ChilOdr<lpur MahMaslrol lndia Sakrebail Karnoltaka - 2 batches India Vimoli Vir. Kamalolkiln - 2boltches India Karadibella Kamat<lka

India Gilillegundi Kamat<lkil India Maukal Kamataka - 7IMtches

Thail.lndMoileHualloilmpang (nalural sland) Thaîland~iat'Huai loiImpang (planled stand)

BuluKumba, Il1dont'Sia

Papw New Guinea ex Brown Ri\·er· presumably {rom Myanmilr (Bumw) (Cameron1966,White, personal communication);

SalomonIsbnd Arara - prt'Sumably introduœdfromMyanmarif notfrom India orThaiJand, via Papua New Guinea (Whitt'. personal communication);

SalomonIsbnd Vîru-presumably introduced from Myarunarifnot from

IndiaorThailand, via PapuaNewGuinea (White,personal

communication);

Perlis. Peninsular Malaysia - from Forest Research Inslitute Malapia. preswnably (rom Thaibnd

Kota Marudu, Sabah - introduced by the Outch Tobacco Company, presumably fromIndia

Tot.l: 22~edlots ofprogeny and provenances induded, constitule a

real problem; information, when reliable, is fragmentaryandalotof unœrlaintiesremain. In this respect,wcarcpldcîng strong hopes on the use of nlolccul.lr nl,ukers for ,1ITessîng the information that

IS1.1Cklllg.

CERMPlASM CONSERVATION

The teak genetic resources gathered within the projett canbe conservee! either as seeds, planted outdoors or in tissue cultureconditions.

Asseells

Seee! lots canbe stored in a fridge or cold room, at

4°C for sorne lime. However, 10ss of germination capacity over time must nol be undereslimated. Several months of storage under such environmenl may result in a dramatic dccrease in germination rate, already known 10 be 10w and unpred.ictable under the best conditions for teak (White, 1991). Dcpending on the challenge, resorting to tissue allture for germinating reca1cilrant seeds can be

cnvisaged, as already successfully undertaken (Monteuuisl'tal" 1998).

ln vivo

Wîthin our project,ln VIVOconservation plotsronsist

mainly in the resources existing in the nursery or in the base and broeding populations, in other words, in demonstration plots, provenance-seed source/ progeny trials, clonai tests and seed stands. Practically, mainly gcnotypes exhibiting superior phenotypes will bepreservee!. In that sense, our concept of germplasm conservation, deliberately operation-oriented, might be too restrictive. Cost, however, remains (orusa majorconœm. Wecannot afford to set up and maintain ex-situ conservation plots for ail the genotypcs (rom as many different origins as wc can gcl, owing 10 variotls factors such asavailability of sites, costs in silvicultural practices and manpower for maintenance of the plots

brvitro

ln vitroculture conditions can be agoodoption for

germplasm conservation (Haines, 1994), in a more restricted environment than lnVlOO plots whereby proper maintenance remainsa critical issue(ZobeIand.

T,Ibert,I984).1kp'thog<!n-free_oftis5ue cultureallow the intemational exchange o( living plant

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material without any sanitary or climatic constraints, contrary toex-vitroplants or plant portions.

Table 2. List of Families obtained from the Ivory Coast Clonai Seed Orchard and planted wilhin our project

Currently, our in vitro gene pool is limited to the genotypes under micropropagation. The protocols developed havebeenconœived as very conservative for maintaining the various genotypes for several years under sustainable sub-culture regimes, while preventingsomaclonal variation risks (Monteuuiset

al.,1998; Goh and Monteuuis, 2001).

Clonai or genotypic fidelîty, whieh is a requisite for gene bank, remains a crucial conœm for us. Another option under investigation currently is the resort to cryopreservation, Ihe efficieney of which has been proven for other tree species (Ashmore, 1997). Developing protocols adapted 10 teak will be facilitated by the possibility to regenerate teak plant

fromin vifromeristemculture, contrary to many tree

speeies (Monteuuis, unpublished results). Themain advantage associated with cryopreservation is the possibility to store genotypes for unlimiled periods of times ina very resITicted envirorunent and wîthout maintenance requirements, while preserving their integrity, as detailed by Ashmore (1997).

Mass selection prevails at the firsÎ" step of the selection process. Individual selection is mainly phenotypic, with special attention to traits of major economical importance for teak such as: growth rate, small nodes, baie straightness and basal circularity in absence of butlress or flules. Wood qua lity assessment including bath aesthetic (wood pattern, lexture) and technologiea! characteristic:s has also 10

hetaken intoconsideralion for phenotypic selection.

SEED STAND OPTION

STRATEGIES fOR SEITINC-UP IMPROVED

YIELD AND QUALITY PLANTATIONS.

From seeds

Thedifferent origins listed (fables 1and 2) have been set up on easily accessible ICSB stands according 10 planting designs adapted to the conversion of the demonstration plots or provenances-seed sourcel progeny trial intoseed stands. The Iwo provenancel progeny trials were set up in a partially cquilibrated incomplete block design and were comprised of 41

and42 seedlots respedively, with 26 seedlots rommon tobothtrials(WiIliamsandMatheson, 1996). Thetwo sites were very dissimilar in terrain in lhat one ison a' hilly area (Luasong Forestry Center) whereas the othee is located on lowland (faliwas, Lahad Datu). Data from these trials will thereby also allow us to assess thegeneticoriginxenvirorunent interaction. Further, as much as possible,closevictnityofgenetically related individuals within eath trial has been avoided. Each progeny elementary plot did not indude more than 5 sib1ingsinorcier tominirnizethe lasses when selecting the best in the plot tohekept asseedproducers, while fellingthe others inorder to prevent risks of înbreeding depression.

Thetraditional meansof propagaling leak is through seeds, as has been practieed for centuries, with lhe possibility of storing the seedlings in lhe form of "stumps" when necessary (Kaosa-ard, 1986;Tin Tun, 2000). Various seed-based strategies have been suggesled by specialists for improving the genetie quality of teak seeds (Wellendorf and Kaosa-ard, 1988; Kaosa-ard, 1998, 1999; Kjaer

et

Q/.,2000). The seed stand and the seed orchard options are worth considering in ourcontext. 13 10 3 2 3 2 1 2 1 3 3 3 3 2 2 2 1 Number of seedlots Talai: 56 seedlots 17 origins India Nellicutha India Nilambur India Vemolirge tndia Va India Purunakote Ivory Coast Bamoro Ivory Coast Kokondekro Laos Paklay Senegal Djibelor Tanzania Kihuhwi Tanzania Mt Tanzania Bigwa Thailand Huoi-Nam-Qon Thailand Maasale Valley Thailand Pong Salee ThailandBanPha Lay Thailand Ban Cham Pui

Total:

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ln this respect. wc arc placing a lot of emphasis on theuhlizalion of non-<:!estruclivc methods forwood qualily analysis (Baillères and Durand, 2000). Furtner 10 thiSsclectionof plus tœes fordonaltesting. inferior trees and half-sib in close vicinity are culled

inorder to lavor the intennatingofsuperior indi\idual not gmehcally relaled. In the absence of sufficiently detailed records, rcsorl 10 molecular biology

techniques such as AFLP dc\'clopedinCirad-Forêt Iaboralory canhelpinthedetenninatÎonofthegenetic

relaledness among îndividuals withîn the same

ooghborllood.Thelurtt....testingofthesclectedsœd

produœr.;01 thesœdstmd 00sed on theperlonnanœs of thcir progeny will indicale lheirrombiningability

which can beused for rcfined or more advanœd sclectooandcullingactivilies(lqaerandFaster,1~; Mandai and Chawhaan. 1999). A final densityof 120 - 180seed produœrs per heclare of seed stand îs expected at lhe end of lhese selection and roguing actîvilîes(Figure 1).

SEED ORCHARD OPTION

The besl combining genolypes or "combiners" can he asexually propagatcd or duplicated 10 be mixed

according to a well-suitcd plantmg design within a \-egelalive seed orchard consisting clones from different familiesand provenances, as illustrated in Figure1.

Thesc clones will be produœd on their own root systemaseither rootcd cuttings. microcullings or even layerîng (Lahirî.1985; Monlcuuis,1al , 1995;

Monteuuisria/., 1998)in order to prevcnt grafting mcompalibility problems and Ihe consequenlial production of "ilIegitimates". Theseare likely 10 dcpreciate the genctîc quality of the seeds produœd. firstly, whencollected directly from suffiunexpeeled 'mothers'sinœiIIegitimates aremostofthelimehard

10 dîslinguîsh from the grafts as Ihey look similar. and secondly. as these will pellute the genetîc quality of theseeds produœd by lhe "Iegitimates" around. However, the numerous question marks and unœrtaintiesassocialed with lhe realbenefits thal can beexpected from such orchardshave 10 beserîously pondered (Kjaer and Foster. 19%; Kasoa-ardrial.. 1998; Kjaerrial.•2(00;While and Gavinlcrtvatana. 1999).The implcmentation of trus strategyistherefore only 10 beconsidered forthe long lerm.

IMPROVED QUALITY SEED PRODUCTION STRATEGY

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BasePopula'.on (P'o-.enanœ 'po-ogenylnab)

Figull' ): Illustration of the project striltegy aiming al producing improved qualityseedsinthe shorlest delay slartîng from genetic.lll)'-rich base populations. Ad\'anœd generations of clonai seed orchArds canheenvîsaged for the I\ln~lerm

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FROM CUlTINGS OR MICROCUlTlNGS

Production of teak planting stockbyseeds,although primarily cmployed. remains severely handicapped

bythe followings factors:

1. Quantilalively limited seed production (Wellendorf and Kaosa-ard. 1988, White,

1991 ).

IL laie flowering.Ilis noteworthy that in teak,

straight bole length, which affedsdirectlyils

market value.isstrictly dependcnt on the capadty oftheterminal meristem 10 remain vegetative as long as possible (White. 1991). Ils conversion iota the f10wering stage

induœs a lork formation, as the result of a

true dichotomy proœss.

iü. Lowgermination rates (Kaosa-ardttal.,1998; White, 1991). InThailand,for e.xample,onIy

5 plantable seedlings on average can be

p<actically

"'J"'CIed

(romlOOseeds ID'

Luge-scale nursery (Kjaer and Fosier, 1996;

Kaosa-ard,I998).

iv. Substantial variability among individuals,

even amonghalf-sibs,withinprogenie<>,with

regard 10 traits of major economical importance such as growth, form, technological and aesthetic characteristics (wood pallem)(Dupuy andVerl>aegen. 1993; Kaosa-ardetal.,1998; Bath,2<XXI);

v. Limited accurate genetic knowledge about the inheritance of such economically significant traits, and consequently, sorne uncertainty for the ultimate gain, notwithstanding the time conslraints associated with sound breeding programs (Wellendorf and Kaosa-ard, 1988; Kjaer and Foster, 1996; Kaosa-ard, 1998).

The hindranœsand uncertaintiesassociated with long tenn breeding strategies for teak greatly

penalized

by low seedling pnxiuctivity havebeendeve10ped(Kjaer etal., 20lXl; White and Gavinlertvatana, 1999). According to lheseauthors,lhemagnitude ofthereal genetic gain associated withtheseedling route has yet to be c1early defined, and the basic question to know whether ail the efforts invested during the past deeades are worthwhile remains. Thisisundoubtedly

a majorconœm for potential investors, forwhich rapid pay-off is a decisive argument. From practical and theorelical infonnation, il can beob;ectively assumed that for leak, greatergeneticgaincan beexpectcd from the cuHing foreslry option, especially whcn clonai, !han !ha,fromtheseedling10"",')'(ZOOeI.ndT.1bert,

1984; Ahuja and Libby, 19933 and b; Monteuuisand Goh,I999).

Two different strategies can he considered for establishing wood production populations from vegetatively mass-produced teak planting stock, either in bulk fonn or by clonai propagation.

TIlE BULK OrnON

Bulk propagation consists in vegelalively propagaling a group of mixed genolypes withoul mainlainingany individual identification. Thiscan

be useful for increasing the number of a limiled quantity of juvenile genotypes of presumablyhigh

and.sirnilargeneticvalue, derived for example,from controUed pollination.

The main advantage of the bulk propagation option lies in the absence of aneedto strictly and c1early idenlify by proper Jabeling one genolype from another. This option will also maintain a certain degree of genelic variability depending on the numberofgenotypes involved at the beginning, and which in tum may inducean overall heterogeneity in the resulling wood production populations. This seems espe<:ially true for teak considering the variability among genotypes. However, successive generations of seriai propagation may eventually result in a significant reduction of the original genetic base due to genolypic differences in the multiplication rates, i.e. in the number of shoots produœd to beusedascuHings withsuffidentlyhigh ability for adventilious rooting.

Bulk propagation has been reslricted in our project to superior quality, quantitatively limited or rare seed lots propagated under invitroconditions. At lhis stage, the genotypes are too young 10 express any individual differenœ wananting a selection. The original geneticbasein optimized micropropagation conditions, contrary to nursery environment, is thllS maintained at1eastduring the first several subcuJture

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cycles. Howevcr, over lime, with littleor no conlrol on lhe gcnolypc-dependenl capacity for axillary shoot production, the risk of gradually losing clones withrcduœdcapacity hasto beacknowledged.

THE CLONAL OPTION

Talbcrt. 1984; Ahuja and Libby. 1993a and b). The possibilily10reproduce. lheoretically in unlimited numhers. the best Irces. from quality and yield standpoints, for large-scale plantation uses. offers tremcndous prospects.

ln clonai propagation. contrary to the bulk propagation strategy. the genotypic identity is rigorously and individually preserved through successive propagation c)'des. which may last 5e\'eral œnturiesIIIœrtaincases. Eachcloneronsists of a.sexually-derived offspring with virtually the same genetic make-up. rcgardless of the numberof ils represcntatives.

Oonal propagation im'olvingscrupulousgenotypic identification ensures a better control of the plant material propagatedbycuUings than by the bulk option. in addition toa numberof olher advantages associated with doning of foresttrees (Zobel and

The basic requisite for sucœssfultcak propagation by rooted cullings is the exi:.tcnœof agoodcapacity for adventilious rooting (Monlelluisri0/.• 1995).

Fromour first field observations. it appears lhat once rooted. teak cullings de\'elop Htrue-to--I)'J)(''', and a good wilhin-done uniformily can he ultimately expeeted. Increased yield. higher uniformity for economically important traits suchasgrowth rate. trunk form, straight bole length. wood characteristics. and shorler rotations constitute strong inœntivcs todevelop teak donal plantations (Wellendorf and Kaosa-ard,1988;Bath.200).50ch unifonnity can not objectively bcexpected from plantations set up from seedlings, or even from cuUings issucd from bulk propagation by virtueof

CLONAL STRATEGY

Figure2;Illustrationofthepro;ectstrategy forIarge-scalecml! plantations olll.'.ll MolecuL.umarketscan hdpwith l''-'dJgree information,certîfK'atÎQnas weU as for genetk reLlledness between thedonesusedforc\on.lldeploymenl

Advancedsoedioll

Clonai perfonnances IcharacleristlCS acCOl'ding 10 site conditions Additional'NOod(desruclt-/e) analyses Genetic relatedness (rmlecUlar markers)

~D-_~Mlmo_

Mass selection of .... Trec:s ..

basedonphenotypic criteria Famly. 'tNithin family selection Non-deshuctlve wood analysis

methods BasePopulation (Provenance/progeny lrials)

(10)

the arguments devcloped previously. Properly selected and wisely dcployed clones will thus maximize the shorHerm returns From suitable planting sites, which are dramatically reduang in surface area(Balidol" 2(0)).

Utilizalion of clones can he profitably adapled 10 inlercropping after proper selection particularly, on crown form. Planling densily and silviculture practiœs can he adapted ta inlensive managemenl systems. with the p<xssibility to harvestseverallimes

from the same slump, laking advantage of the excellent coppicingability (Martinttal.,2(0)).Soch praclices look very attractive for enhancing plantation yield while significanUy reducing the planling and lhe (micro)cutting rosts. With regard to these financial aspects, ithastaheemphasized thatclonalplantations gener.illyrequirele;,;planting slock than plantations eslablishedfromseedlings, whichcompensate, la a certain extmt.thehighercœt

of (micro)cuttings. Figure2summarizesthedonal stralegy adopled for our project:.

Elficitnttechlliquu10rmassfJt!gt!latroeproduction

01silpaiorquality planting stock

Massproduction

of

rootnJcuttings in nursny

Mass propagation of leak of any age by rooled cultings in nursel)' conditionshashemdeveloped

andbecome fully operational within our projectsinœ

1992.Therequisites, as weil as the advantages and Iimils of Ihis propagation option have been exlensively prescnted and confirmed by a decade of experience (Monteuuis, 1995; Monteuuisd 01., 1995; Monleuuis, 2(0)). Average rooting rates of 8O%are routinely oblained From malure genotypes intensively managed as container-grown stock plants once the mobilizalion phase has been sucœssfully carried out. In such conditions, annual production of600 rooted cuttings per square meterof stock plant area (15 stock plants per square meler) q1n he easily oblained (Montcuuis el al., 1995).

lnvitro mass propagation

Theavailabilityofa well-equipped Iaboralol)'within our jointpro;eetprompted us to explore the prospects of propagaling teak in lissue rulture conditionsvia

microcuttings (Bon and Monteuuis, 1996). This suceessful application has already becn reported (Monteuuisetal., 1998;Goh and Monteuuis,2IXlI).The conœivedtissueculture protocolswcremadeas simple as possibleinorder 10 beeasily applicable and tocope with the constraints of large-scale application in tenns of cosl efficiency and high produclivity. Mass micropropagation of any genotype, eitherinbulk or cIonaUy,throughaxill<uy-produœdrrticrœhootswith

anexponential multiplication rate of three to four at lheend off!Verysixweek-duration sub<ulturesis now possible. The rooting-acclimatization phase is advanlageouslyachieved in nurseryconditions under a misI-system with more than

m

SlJ<.'ŒS501laverage.

Afterweaningand

raisins

onder thesame intensive nursery conditions as for the cutting.œnved plants, thetissue-cu1ture plantsdevelopiolo vigorousand

true-to-type vegetalive plants. More than

400.00>

miaocu~havebeen produœd up tonowforkx:al

planlationsas weil as for oversea markets by applying

!lUstedm;que.ThepœsibililyIDsend thein"",,,plants offta differenl destinations, regardlessofdistance, in the absence of phyto-sanitation restrictions, considerably expands lhe market prospects and constitutesanoutstandingasset.

The invioorooted cutting and invitromicrocutting options forvegetatively mass propagating superior teak genotypes with the respective pros and cons have been extensively reviewed, leading to the conclusion that the best oplion, in many respects, consîsts in the combination of the Iwo techniques (Monteuuis,2OO)).

CONCLUSION AND PROSPECTS

The Research and Development collaborativeproject belween less and Cirad-Forêt was inilially implemented towork primarilyon rattanspecies. The shift and the emphasis given to teak started in the carly 1990's as a result of the unexpected sucœss obtained

fromthevegetative

propagation

lechniquesdeveloped at the nursery and tissue culturelevels. The possibility 10 mass produœ clones by rooted cuttingsor invitro microcuttings in cost-effective conditions from any selected mature superior teak

tree,

as weil as from good-performing teak Irees introduced locally, prompted us to inves! more ontheseadivities. The rationaleandthe prospects of using clonal materia1s

(11)

for leak have been indœd advOGlled for a long lime. Beingawareofour vegetative propagationassets,the nexlstep!las been10gatherour ownbasepopulations

....Tilhagcneticbackground asrichaspossible Thisaim !lasbeen attained to a large exlenl by our possesslon of a highly diverse teak genetic resourœ thal covers Ihe range of adaplabilily of Ihe spedes. This noleworthy genetic richness combined 10efficienl propagation and diffusion lechniques are Iikely10

satisfy any plant malerial arder,intheform ofseeds,

or clones in tissueculture conditions for over5ea markets, with Ihe possibilily 10œrtify Ihe plant material byDNAfingerprinting.

Keeping up wilh the Ialest advances in lechnology which can further bcnefit lhe projet! rcmainsa major conccm. In this respect, we are placing emphasison the application of motecular markers such as microsatellites and AFLP lechniques currently cmployed for leak by Cirad-Forêt, as weil

as

non-deslructive woocl analysis mclhods for moving one slep forward in the selection of superior pIani malerial and clonai dcployment. Proper sitex genotype matching indeed deserves particular attention withrespecttopestand disease(defoliators and borers) aspects, as weil as to heartwood formation, controlledbygenetîc and environmental parameters, and which delennînes log qualily (Bailleresand Durand, 2OOl}

Theneed to intensify the production of premium quaütylùghv.>Iue tin>œs..eakbeingthemostprized

one, with a worldwide demand far grea!er lhanthe supply available, isa slrong inœntivealong this Hne.

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